<p>Locomotion in any environment requires the use of energy to overcome the physical</p><p>forces inherent in the environment. Most large marine vertebrates have evolved</p><p>streamlined fusiform body shapes to minimize the resistive force of drag when in</p><p>a neutral position, but nearly all behaviors result in some increase in that force.</p><p>Too much energy devoted to locomotion may reduce the available surplus necessary</p><p>for population-level factors such as reproduction. The population of North Atlantic</p><p>right whales has not recovered following legal protection due to decreased fecundity,</p><p>including an increase in the intercalf interval, an increase in the years to first calf and</p><p>an increase in the number of nulliparous females in the population. This reproductive</p><p>impairment appears to be related to deficiencies in storing enough energy to meet the</p><p>costs of reproduction. The goal of this study was to determine whether increases in</p><p>moving between prey patches at the cost of decreased foraging opportunities could</p><p>shift these whales into a situation of negative energy gain. The first step is to</p><p>understand the locomotor costs for this species for the key behaviors of traveling and</p><p>foraging.</p><p>This study investigated the cost of locomotion in right whales by recording the</p><p>submerged diving behaviors of free-ranging individuals in both their foraging habitat</p><p>in the Bay of Fundy and their calving grounds in the South Atlantic Bight with a</p><p>suction-cupped archival tag. The data from the tags were used to quantify the oc-</p><p>currence of different behaviors and their associated swimming behaviors and explore</p><p>three behavioral strategies that reduce locomotor costs. First, the influence that</p><p>changes in blubber thickness has on the buoyancy of these whales was investigated</p><p>by comparing the descent and ascent glide durations of individual whales with differ-</p><p>ent blubber thicknesses. Next, the depth of surface dives made by animals of different</p><p>sizes was related to the depth where additional wave drag is generated. Finally, the</p><p>use of intermittent locomotion during foraging was investigated to understand how</p><p>much energy is saved by using this gait. The final piece in this study was to deter-</p><p>mine the drag related to traveling and foraging behaviors from glides recorded by</p><p>the tags and from two different numerical simulations of flow around whales. One, a</p><p>custom developed algorithm for multiphase flow, was used to determine the relative</p><p>drag, while a second commercial package was used to determine the absolute mag-</p><p>nitude of the drag force on the simplest model, the traveling animal. The resulting</p><p>drag estimates were then used in a series of theoretical models that estimated the</p><p>energetic profit remaining after shifts in the occurrence of traveling and searching</p><p>behaviors.</p><p>The diving behavior of right whales can be classified into three stereotyped be-</p><p>haviors that are characterized by differences in the time spent in different parts of the</p><p>water column. The time budgets and swimming movements during these behaviors</p><p>matched those in other species, enabling the dive shapes to be classified as foraging,</p><p>searching and traveling behaviors. Right whales with thicker blubber layers were</p><p>found to perform longer ascent glides and shorter descent glides than those with</p><p>thinner blubber layers, consistent with the hypothesis that positive buoyancy does</p><p>influence their vertical diving behavior. During horizontal traveling, whales made</p><p>shallow dives to depths that were slightly deeper than those that would cause ad-</p><p>ditional costs due to wave drag. These dives appear to allow whales to both avoid</p><p>the costs of diving as well as the costs of swimming near the surface. Next, whales</p><p>were found to glide for 12% of the bottom phases of their foraging dives, and the</p><p>use of `stroke-glide' swimming did not prolong foraging duration from that used by</p><p>continuous swimmers. Drag coefficients estimated from these glides had an average</p><p>of 0.014 during foraging dives and 0.0052 during traveling, values which fall in the</p><p>range of those reported for other marine mammals. One numerical simulation deter-</p><p>mined drag forces to be comparable, while the other drastically underestimated the</p><p>drag of all behaviors. Finally, alterations to the behavioral budgets of these animals</p><p>demonstrated their cost of locomotion constitutes a small portion (8-12%) of the</p><p>total energy consumed and only extreme increases in traveling time could result in a</p><p>negative energy balance. In summary, these results show that locomotor costs are no</p><p>more expensive in this species than those of other cetaceans and that when removed</p><p>from all the other stressors on this population, these whales are not on an energetic</p><p>`knife edge'.</p> / Dissertation

‘Critical Habitat’ is the habitat required to close the life history of an endangered species and is a fundamental requirement for species recovery for two reasons; the role of habitat in population limitation and viability must be determined, and the habitat must be protected. The North Atlantic right whale is an endangered species that annually migrates to the Grand Manan Basin and Roseway Basin Critical Habitats to feed on diapausing calanoid copepods that are typically aggregated at depths of 100 to 150 m. In this thesis I quantify spatial and temporal variation in the copepod prey field and occupancy of right whales in Roseway Basin, and use this information to identify the location and extent of right whale Critical Habitat. To accomplish this, I measured copepod abundance and energy density (kJ m-3) using optical, acoustic and net collection methods during 2007 to 2009. Oceanographic processes that affect variation in the copepod prey field include slope water intrusions, water mass density, gyre-like circulation and frontal features. Aggregations of diapausing copepods are maintained on the southern slope of Roseway Basin by cross-isobath tidal advection, and are advected along-isobath by the residual flow. Tidal advection at a front, coupled with along-isobath advection and shear in the horizontal currents serve to accumulate copepods along the slope where aggregations are maintained for at least 7 days. The abundance, stage-structure, species composition and aggregation locations of copepods, as well as the hydrography and circulation, were variable among the three years of the study. A 20 year time series of right whales, copepods and hydrography revealed that interannual whale occupancy in the Critical Habitats is variable and can be explained by prey field variation only in Roseway Basin. Factors other than the local prey field affect the number of whales that occupy Grand Manan Basin. Variation in the right whale prey field could not be explained by temperature and phytoplankton-dependent growth in the Scotia - Fundy -Gulf of Maine region. The results of this thesis assisted in establishing the Roseway Basin right whale Critical Habitat in 2008, and the cross-disciplinary nature of the study also provides new insights into the relationships between biology and physics in Scotian Shelf - Gulf of Maine basins.